ASIAN MYRMECOLOGY Volume 6, 63–82, 2014 Issn 1985-1944 © Ba d a m d o r j Bayartogtokh , Ul y k p a n Ai b e k , Se i k i Ya m a n e a n d Ma r t i n Pf e i f f e r
Diversity and biogeography of ants in Mongolia (Hymenoptera: Formicidae)
Ba d a m d o r j Bayartogtokh 1, Ul y k p a n Ai b e k 2, Se i k i Ya m a n e 3 a n d Ma r t i n Pf e i f f e r 2
1Department of Zoology, School of Biology and Biotechnology, National University of Mongolia, Ulaanbaatar 210646, Mongolia 2Department of Ecology, School of Biology and Biotechnology, National University of Mongolia, Ulaanbaatar 210646, Mongolia 3Graduate School of Science and Engineering, Kagoshima University, Kagoshima, 890-0065 Japan
Corresponding author’s e-mail: [email protected]
ABSTRACT. Information on the biogeographical composition and spatial distribution patterns of ant assemblages in Mongolia is relatively scarce. In this study we investigated species richness, spatial distribution and biogeographical composition of the Mongolian ant fauna and recorded a total of 71 species belonging to 17 genera and three subfamilies. Genus and species richness of ants is lower in Mongolia than in most surrounding geopolitical regions. Using published literature as well as our own data we examined how ant species richness was related to phytogeographical elements. The major part of the Mongolian ant fauna is associated with the mountain taiga and forest-steppe ecosystems. A few ant species belong to the arid steppe, semi-desert and desert ecosystems. We observed no significant correlation between species richness of ants and land area size of phytogeographical regions; however, species richness of ants and plants in various phytogeographical regions showed a significant positive correlation. We did not detect strong differentiation of ant species along the elevational gradients of the country as most of the species were recorded in a wide range of elevations, but mid-elevational altitudes (1000 – 2000 m a.s.l.) showed the greatest diversity of ants. Regarding the biogeographical composition, species with wide geographical distributions, i.e. Transpalaearctic species together with Eastern Palaearctic, Central and East Asian elements, comprise a large proportion of the ant fauna in Mongolia. The ant fauna of Mongolia does not differ significantly from that of neighbouring regions, and the majority of species occurring in this country probably originated in neighbouring areas to the west, north and east.
Keywords: Formicidae, area size, biogeography, distribution, diversity, elevation, Mongolia
INTRODUCTION Mocsáry & Szépligeti (1901) and Ruzsky (1903, 1915) published the pioneer works on Ants are common, almost ubiquitous insects in the Mongolian ants, based on material from Mongolia, and they can be found in all ecosystems the southern Gobi and northern forested areas. and main terrestrial habitats. However, the ant However, the exact locality of some historical fauna of Mongolia is still incompletely known. records from southern and northern Mongolia is
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unclear, because these places are now in China genera, with new records of six species. They or Russia, and difficult to access. For instance, excluded several species from the checklist due to Holgersen (1943) recorded Myrmica rubra improbable occurrence or incorrect identification. (Linnaeus, 1758) from the place named as Sajan, Subsequently, Yamane & Aibek (2007) published Sistikem, but Wetterer & Radchenko (2011) a short note on ants of Mongolia that included all revealed that the actual name of this place is species also included in Pfeiffer et al. (2007). Systyg Khem of the Sayan Mountains (in Siberia), Later studies focused on certain which is located in the Russian Republic of Tuva. systematic groups, rather than on documentation Another example is Stitz (1934), who reported of overall faunal diversity. Thus, Aibek & five species, namely Formica clara Forel, Yamane (2009) reviewed the genus Camponotus 1886, F. picea Nylander, 1846, Lasius alienus in Mongolia, and recorded five species along with (Foerster 1850), Tapinoma geei Wheeler, 1927 a new finding of two species,C. aterrimus Emery, and T. orthocephalum Stitz, 1934 from south and 1895 and C. (Tanaemyrmex) tashcumiri Tarbinsky, southwestern Mongolia, without giving exact 1976. Subsequently, these authors studied the names of the collection localities. Two of the subgenera Austrolasius and Dendrolasius of the above-mentioned species, T. geei and L. alienus, genus Lasius in Mongolia (Aibek & Yamane have not yet been confirmed in Mongolia. It is 2010), and reported nine species of Lasius, two possible that the collection sites are located in of which, namely L. (Austrolasius) reginae Faber, Inner Mongolia, China. 1967 and L. (Dendrolasius) fuji Radchenko, Actual research on ants in Mongolia is 2005, were new records for the Mongolian fauna. considered to have commenced in the mid 1960s Lasius obscuratus Stitz, 1930, which had been with Dlussky (Dlussky 1965). Later, Pisarski excluded from the Mongolian fauna by Pfeiffer (1969a, b), Dlussky & Pisarski (1970), Pisarski et al. (2007), who doubted the occurrence of this & Krzysztofiak (1981) and Radchenko (1994a) species in Mongolia, was again listed by Aibek & published extensive works and recorded more Yamane (2010). However, the nominal subgenus than 50 species for the country. However, this Lasius has not been seriously revised, leaving number required some amending to reflect the many problems. taxonomic status and actual occurrence of some More recently, Yamane & Aibek (2012) species in Mongolia. Therefore, Radchenko studied the distributional patterns of 14 species (1994a, b, 1995, 1997, 2005) and Seifert (2000, of the genus Myrmica in Mongolia, and provided 2002, 2004) provided additional information on an identification key for all those species. the ants of Mongolia, and were concerned with the They omitted Myrmica commarginata Ruzsky, taxonomic resolution of the species. Pfeiffer et al. 1905 from the species list, but newly added M. (2003) completed the first comprehensive study lobicornis Nylander, 1846. on ant communities in Mongolia examining the A significant focus for present-day distribution patterns of ants along an ecological myrmecologists is the assessment of biodiversity, gradient from steppe to desert, assessing the community composition, biogeography, and other impact of climatic parameters and the influence basic investigations of the ecology of a regional of vegetation on community structure and species ant biota. Such a study would grant researchers diversity of ants. Later, Aibek et al. (2006) a background from which to begin further reported 17 species from Mt Bogdkhan in north- more detailed studies. The research presented central Mongolia along with information on here examines the ant fauna of Mongolia in a nesting habits of each species. They reviewed the biogeographical and ecological context. For the ant species list recorded in this mountain area, moment we simply attempt to combine what is and made comments on status and earlier records already known with our unpublished data in order of some of those species. to contribute to a better understanding of the summarising the literature sources as diversity and biogeography of ants in Mongolia. well as their unpublished records, a checklist of Hopefully, this work will also help to conserve ants in Mongolia was published by Pfeiffer et the interesting ant fauna of Mongolia. al. (2007), which reported 68 species from 17
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Fig. 1. Phytogeographical regions of Mongolia (modified after Yunatov 1950). Refer to “Study area” subsection for numbering of these regions. White dots represent major cities, black dots the sampling points of our study. Dark green represents mountain taiga forest, light green – forest steppe, yellowish green – steppe, yellow – semi- desert, tawny – desert ecosystems.
& Dagvadorj 2010). The highest precipitation MATERIALS AND METHODS values do not exceed 600 mm in the northern mountains, whereas southern desert areas receive Study area roughly 50 mm of precipitation. From the north to the south of the country, mean annual precipitation Mongolia stretches across central and north- decreases steadily, which is a decisive factor for east Asia, and occupies several vegetation zones the distribution of vegetation formations. From where the Siberian taiga forest meets the Central north to south it not only gets drier, but warmer Asian steppe and desert. The north-south axis as well. From the west to the east some areas measures more than 1,200 km at its maximum, show comparatively low precipitation because and the widest east-west distance reaches about they are on the lee side of mountain chains. Such 2,370 km. Mongolia is an upland country, with an a rain shadow effect is most pronounced in the average elevation of 1,580 m (range of elevation: depressions of larger lakes in western Mongolia, 560 – 4374 m), and about 85% of its area elevated but it is recognisable in central and eastern over 1,000 m a.s.l. (Murzaev 1952). Mongolia (Hilbig 1995). This complex set of Mongolia has an extreme continental parameters, together with the large area of the climate, and temperature fluctuates greatly, both country, creates a large variety of habitats and daily and annually. As estimated for about 60 years has a distinct influence on the biodiversity of the between 1940 and 2000, the lowest temperatures region. are recorded in January, with monthly averages Temperature and vegetation follow under −15°C and minimum temperatures as low a similar pattern of altitudinal zonation, and as −30°C. July is the warmest month, with mean generally speaking, the north to south distribution temperature 15°C in the mountains and 25°C in of vegetation zones corresponds well with the the southern semi-desert and deserts (Natsagdorj precipitation pattern. Thus, Mongolia has several
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vegetation zones and vertical belts, i.e. from org; http://www.antbase.net; http://www.antcat. north to south we meet: alpine meadow (alpine org; http://antwiki.org). In the present paper, the vegetation with upper limit of closed turf), species list by Pfeiffer et al. (2007) is updated mountain taiga-forest, forest-steppe, steppe, with some modifications. Generic and species desert-steppe (semi-desert), and desert (Yunatov names of ants are listed in alphabetical order. 1950; Murzaev 1952). The relative area of the phytogeographical Based upon vegetation, floral regions of Mongolia is taken from Ulziikhutag composition, topographic and climatic (1989). Plant species richness was estimated characteristics, Mongolia divides into 16 based on the number of vascular plant species phytogeographical regions (Yunatov 1950; recorded within each phytogeographical region Murzaev 1952; Grubov 1955, 1982; Fig. 1). (Gubanov 1996; Ariuntsetseg & Boldgiv 2009). Two of these regions are mountain taiga forests Biogeographical analysis of ant species (1. Khuvsgul, 2. Khentii), three are forest- found in Mongolia is based on data in the steppes (3. Khangai, 4. Mongolian Dauria, 5. published literature which provides information on Great Khyangan), four are steppes (6. Khovd, 7. geographical ranges (Seifert 1992, 2000; Bolton Mongolian Altai, 8. Middle Khalkh, 9. Eastern 1995; Kupyanskaya 1995; Collingwood 1997; Mongolia), four are semi-deserts (10. Depression Czechowski et al. 2002; Bolton 2003; Imai et al. of Great Lakes, 11. Valley of Lakes, 12. Eastern 2003; Radchenko 2005; Aibek & Yamane 2009, Gobi, 13. Gobi Altai), and three are desert regions 2010; Radchenko & Elmes 2010; Guénard & (14. Dzungarian Gobi, 15. Trans-Altai Gobi, 16. Dunn 2012). Division of biogeographical regions Alashan Gobi). is based on that applied by Bielawski (1984), Nikolajev& Puntsagdulam (1984), Bayartogtokh Data collection (2010) and Bayartogtokh et al. (2012).
The material studied has been obtained from our Data analysis collections as well as from samples provided by other zoologists and students, derived from We assessed the similarity of the ant species in different regions of Mongolia. Our collections Mongolia and other surrounding regions using
came from 202 sites covering 993 samples, Sørenson-Dice’s qualitative Index (Cs; see
which involved more than 10,500 specimens. Jackson et al. 1989). If Cs= 0, the faunas are
Sampled sites covered the main habitats in all completely different from each other, if Cs= 1, phytogeographical regions of Mongolia though the faunas are identical. Cluster analysis was the number of samples in each region was performed using Euclidian distance measure and different because of their varying area, habitat the complete linkage clustering algorithm with the types and remoteness (see Fig. 1). Ant collections software BioDiversity Professional (McAleece et are currently deposited in the Department of al. 1997). Ecology, National University of Mongolia, The occurrences of ants in different Ulaanbaatar, Mongolia. altitudes and phytogeographical regions were In addition, we collected all previously arranged in presence/absence tables. Multiple available information on ant species in Mongolia, regression analysis was used to examine including revisions of species groups (Mocsáry relationships of ant diversity versus plant diversity & Szépligeti 1901; Ruzsky 1903, 1915; Dlussky and area size. 1965; Pisarski 1969b, a; Dlussky & Pisarski 1970; The differences in the number of plant Pisarski & Krzysztofiak 1981; Radchenko 1994a, and ant species and in the percentage of cover areas 1994b, 1995; Seifert 2000, 2002; Pfeiffer et al. between the various phytogeographical regions, as 2003; Seifert 2004; Radchenko 2005; Aibek et al. well as differences in the number of ant species 2006; Pfeiffer et al. 2007; Yamane & Aibek 2007; along various elevational gradients, were tested Aibek & Yamane 2009, 2010; Seifert & Schultz using one-way analysis of variance (ANOVA). 2009; Yamane & Aibek 2012), as well as species We used Pearson’s product-moment correlations lists from the internet (e.g. http://www.antbase. to examine the relationships of species richness of
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Fig. 2. Relationship of generic and species diversity of ants in Mongolia and its surrounding regions. Abbreviations: JPN – Japan; IRN – Iran; SKO – South Korea; XIC – Xinjiang, China; KYR – Kyrgyzstan; NKO – North Korea; RFE – Russian Far East; URL – Ural region of Russia; MON – Mongolia; WSEK – Western Siberia and eastern Kazakhstan; IMC – Inner Mongolia, China; HJC – Heilongjiang, China; JLC – Jilin, China; GAC – Gansu, China.
ants versus area size, and plant diversity versus ant On the other hand, we omitted Lasius flavus diversity. All statistical analyses were performed (Fabricus, 1781) and Myrmica rubra as these do with software Statistica 5.0 for Windows (StatSoft not occur in Mongolia (Aibek & Yamane 2010; Inc.). For all statistical tests, we considered results Wetterer & Radchenko 2011). We also removed significant whenp < 0.05. Temnothorax serviculus (Ruzsky, 1902) from the fauna list because its former subspecies, T. mongolicus (Pisarski, 1969), exists in Mongolia RESULTS (Pisarski 1969b), and is now an independent species (the nominal subspecies does not occur Diversity of ants in Mongolia). Camponotus japonicus Mayr, 1866 was also removed from the list because its former Based on our investigations, the ant fauna of subspecies, C. aterrimus is now considered an Mongolia comprises 71 species belonging to 17 independent species, while the nominal subspecies genera in three subfamilies. A list of ant species and C. japonicus does not occur in Mongolia (Aibek their occurrences in various phytogeographical & Yamane 2009). Kupyanskaya (2012) has regions of Mongolia is given in Appendix 1. With listed Formica japonica Motschoulsky, 1866 as regard to our species list, the discrepancy with distributed in Mongolia, but its occurrence in this the data from Pfeiffer et al. (2007) who listed country has not been confirmed. So, we omitted 68 species for this country arises because six this species from the list. species (Camponotus aterrimus, C. tashcumiri, Yamane & Aibek (2012) considered Formica orangea Seifert & Schultz, 2009, Lasius the occurrence of Myrmica ruginodis Nylander, reginae, L. fuji and Myrmica lobicornis) have 1846 in Mongolia improbable as this species recently been added to the fauna of Mongolia was not discovered during the recent extensive (Aibek & Yamane 2009; Seifert & Schultz 2009; explorations throughout the country. However, Aibek & Yamane 2010; Yamane & Aibek 2012). we include this species in the checklist based
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on Radchenko & Elmes (2010), who provided a regions of northern China showed much lower distribution map in which the geographical range richness of ants, but this is most probably due of M. ruginodis covered northern Mongolia. to insufficient investigation of those areas (see The largest number of Mongolian Discussion for references and species lists). species belongs to the subfamily Formicinae The ant fauna of Mongolia shares (39 spp.), followed by Myrmicinae (29 spp.); relatively few species with those of the these two subfamilies comprise more than 95 per surrounding regions as a proportion of the cent of the total ant species. A third subfamily, total number of species in compared regions or the Dolichoderinae, is poorly represented in countries; hence the estimated faunal similarity Mongolia with only three species (Appendix 1). was very low. The Russian Far East and Gansu The most species rich and commonly Province of China showed the closest similarity encountered genera were Formica (19 spp.), with the Mongolian ant fauna. Of the countries Myrmica (14 spp.), Lasius (7 spp.), Camponotus examined, Japan and Iran had the lowest faunal (5 spp.) and Proformica (5 spp.). The other genera similarity with Mongolia (Table 1). involved less than five species. Temnothorax and Tetramorium were both represented by Distribution of ants in phytogeographical regions four species, and Leptothorax, Messor and Tapinoma by two species only. Seven genera The number of ant species in the 16 were represented by a single species, namely: phytogeographical regions ranged from 3 to Cardiocondyla, Cataglyphis, Crematogaster, 39 with a mean of 19. Each of these regions Dolichoderus, Harpagoxenus, Plagiolepis and had a peculiar composition of ants, but there Polyergus. Thus, only a few ant genera were were several species dominating in most species rich, whereas the majority comprised regions. Thirty-nine species were found in the fewer species, with the mean number per genus Khentii mountain-taiga region, 38 species in 4.2, and the median only 2.0. the Mongolian Dauria forest-steppe region, 30 We compared the richness of ants in species in the Khangai forest-steppe region, 28 Mongolia with that of the surrounding countries species in the Middle Khalkh steppe region, 25 or regions, and found that both genus and species species in Mongolian Altai mountain-steppe, richness of ants in Mongolia were lower than 24 species in Depression of Great Lakes desert- in most neighbouring regions (Fig. 2). Only the steppe regions, and 20 species in the Eastern
Table 1. Faunal similarity of the Mongolian ant fauna with that of the surrounding regions. Total number of
species, number of overlapping species, and Sørenson-Dice’s Index of similarity (Cs) values.
KYR URL WSEK IRN RFE XIC IMC HJC GAC JLC SKO NKO JPN Number species in 111 76 70 166 79 128 26 32 26 32 136 99 275 each country/ region Number of common 26 24 20 11 25 26 12 12 16 11 20 27 9 species with Mongolia Similarity with 0.22 0.25 0.22 0.08 0.25 0.21 0.20 0.19 0.25 0.18 0.16 0.24 0.05 Mongolian
ant fauna (Cs)
Refer to Fig. 2 for abbreviations of the names of the countries/regions. Only native species are considered, exotic species are excluded.
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Gobi desert-steppe region. The species richness another, and reflected the main landscape pattern of ants in other phytogeographical regions varied of Mongolia. For instance, the boreal forest group between 3 and 18 (see Appendix 1). includes the Khentii, Khangai and Mongolian One species, Formica candida Smith, Dauria regions. The dry steppe group includes 1878, was common to 12 phytogeographical Middle Khalkh, Eastern Mongolia and Mongolian regions, and another species, Proformica Altai regions. The semi-desert group involves mongolica Emery, 1901, inhabited 11 out of Gobi-Altai, Eastern Gobi, Depression of Great 16 regions which provide a variety of habitats. Lakes and Khovd regions. The Gobi desert group Three species, Cataglyphis aenescens (Nylander, encompasses Trans-Altai Gobi, Eastern Gobi, 1849), F. kozlovi Dlussky, 1965 and Tetramorium Dzungarian Gobi, Valley of Lakes and Alashan tsushimae Emery, 1925 were found in ten regions. Gobi regions. The main surprising pattern here Formica lemani Bondroit, 1917 was recorded from was the relatively low dissimilarity observed nine regions, and two other species, F. clarissima between two distant and ecologically different Emery, 1925 and F. pisarskii Dlussky, 1964, regions, such as Khuvsgul mountain taiga region were distributed in eight regions. The other 63 and Khovd mountain-steppe region. We consider species were confined to seven regions or fewer. that this was because only a few wide-ranging From our data it was apparent that 13 species are species (7 spp.) were found in Khuvsgul region. rather rare in Mongolia, their distribution being It should be noted here that because of different restricted to a single phytogeographical region sampling effort, diversity of ants in some regions, (see Appendix 1). A large number of species were such as Khuvsgul, Great Khyangan, Valley of restricted to only two or three regions (12 and 9 Lakes, Dzungarian Gobi, Alashan Gobi etc. might spp., respectively). be not fully explored yet. Overall, the ant faunas Differences between the faunal in various regions were relatively distinct, and this compositions of ants in various phytogeographical fact confirms the well-established classification regions are of great interest. Regions were of the phytogeographical regions in Mongolia. grouped into four main clusters (Fig. 3), and the dissimilarity of the regions reflected their different Species-area relations geographical position and discrepant ecological conditions. Most of these clusters involved the Regarding the geographical distribution of ant regions that are geographically adjacent to one species in various phytogeographical regions of
Fig. 3. A dendrogram depicting the faunal dissimilarity of ants among the phytogeographical regions of Mongolia.
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Mongolia, we found a mismatch between the forest-steppe region that covers only 6.62% of the regions’ extent and the respective number of ant country. Thirty species were recorded in Khangai species. The highest number of species (39 of a total forest-steppe region, which comprises 17.59% of 71 spp.) was recorded in Khentii mountain-taiga the entire area of the country. Species richness region, though the total area of this region is only of ants and total land area of regions were not 3.05% of the territory of Mongolia. Thirty-eight correlated (Pearson’s correlation, n = 16, r = 0.44, species were found from the Mongolian Dauria p = 0.08, n.s., Fig. 4).
Fig. 4. Non-significant relationship between area size and ant diversity in various biogeographical regions of Mongolia. A regression line was fitted to visualise the hypothesised interdependence.
Fig. 5. Relationship between plant and ant diversity in various biogeographical regions of Mongolia. A regression line was fitted to visualise the connection between the two variables.
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Fig. 6. Altitudinal ranges of ants in Mongolia, showing number of species in each category of altitudinal range, with a minimum range of 300 m and a maximum of 1,900 m (size of altitudinal range is given in parentheses).
since area size and plant diversity were 1925, were restricted to lower altitudes, between not correlated for the phytogeographical regions 600 and 800 m a.s.l. Several other species (Formica (Pearson’s correlation, n = 16, r = 0.31, p = 0.24), exsecta Nylander, 1846, Lasius niger Linnaeus, we plotted them separately against ant species 1758, Myrmica lobicornis, Polyergus nigerrimus richness. Within the study region as a whole, Marikovsky, 1963, Proformica kaszabi Dlussky, plant diversity in different phytogeographical 1969, Tetramorium armatum Santschi, 1927, regions varied significantly (ANOVA,df = 1, F = T. concaviceps Bursakov, 1984 and T. inerme 21.16, p = 0.0004). Diversity of ants and vascular Mayr, 1877) were reported at elevations of 600 – plants within phytogeographical regions showed 1600 m a.s.l. Twelve species (Formica candida, a significant positive correlation (Pearson’s F. cunicularia Latreille, 1798, F. kozlovi, F. correlation, n = 16, r = 0.61, p = 0.01) (Fig. 5). lemani, F. manchu Wheeler, 1929, F. pisarskii, F. uralensis Ruzsky, 1895, Leptothorax acervorum Altitudinal pattern of diversity Fabricius, 1793, L. muscorum Nylander, 1846, Myrmica divergens Karavaiev, 1931, M. eidmanni Our calculation of the altitudinal distribution Menozzi, 1930, M. kasczenkoi Ruzsky, 1905) of ant species was based upon collection data were present at the widest range of elevations, from 307 different elevational points. There between 600 and 2,400 m a.s.l. Species richness was no strong differentiation in ant community was greatest between 1,000 and 2,000 m a.s.l., as composition along an altitudinal gradient, as most 27 species were recorded in this mid-elevational of the species were recorded in a wide range of band, whereas 22 species occurred at 800 – 1,800 elevations (Fig. 6). The width of altitudinal range m a.s.l. Thus, we found a peak of species richness was very large for most of the ant species, hence between 800 and 2,000 m a.s.l. the number of species in various altitudes did not differ significantly (ANOVA:df = 1, F = 1.64, p = Geographical distribution of species 0.24). Only four species, Myrmica commarginata, Proformica coriacea Kuznetsov-Ugamsky, 1927, The geographical distributions of all known Tapinoma orthocephalum and T. sinense Emery, species of ants in Mongolia were compiled, and
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the species were divided into groups based upon was described by Pisarski (1963) from Mongolia, their range. Table 2 shows the biogeographical and has not been found outside the country (Bolton composition of the ant fauna of Mongolia. Species 2010). The absence of Oriental or Afrotropical with a wide geographical distribution comprise faunal components in Mongolia is notable. a large proportion of the ant fauna of Mongolia. Three species, Leptothorax acervorum, L. muscorum and Tetramorium tsushimae are widely DISCUSSION distributed throughout the Holarctic Region. The quantitatively most important biogeographical Species richness elements were Transpalaearctic (24 spp., 33.8%), Central Asian (15 spp., 21.1%), Eastern Palaearctic Mongolia has representatives of about 7.4% (13 spp., 18.3%) and East Asian species (10 of the known world ant genera and 0.52% of spp., 14.1%). A few species, Myrmica arnoldii the currently described species (Bolton 2010), Dlussky, 1963, M. commarginata, M. pisarskii altogether 71 species from 17 genera – a relatively Radchenko, 1994 and Polyergus nigerrimus, are impoverished fauna compared to most of the restricted to Siberia and Mongolia (4 spp., 5.6%). surrounding regions. Thus, Tarbinsky (1989) Formica pressilabris Nylander, 1846 is the only found 103 species belonging to 23 genera from representative of European-Siberian species Tien-Shan and Alai mountains in Central Asia (1 sp., 1.4%). There was only a single species, (Kyrgyzstan); Schultz et al. (2006) and Borowiec Harpagoxenus zaisanicus Pisarski, 1963, that et al. (2009) recorded 111 species (22 genera) appears to be endemic to Mongolia. This species from Kyrgyzstan; Gridina (2003) recorded 76
Table 2. Biogeographical composition of ants in Mongolia.
Species Species name Geographical range number Leptothorax acervorum, L. muscorum, Tetramorium tsushimae 3 Holarctic Cataglyphis aenescens, Crematogaster subdentata, Formica 24 Transpalaearctic aquilonia, F. candida, F. clara, F. cunicularia, F. exsecta, F. forsslundi, F. lemani, F. lugubris, F. pisarskii, F. pratensis, F. sanguinea, F. truncorum, F. uralensis, Lasius distinguendus, L. gebaueri, L. niger, L. reginae, Myrmica divergens, M. lobicornis, M. ruginodis, M. sulcinodis, Temnothorax nassonovi Camponotus aterrimus, Formica kozlovi, F. manchu, F. obscuratus, 13 Eastern Palaearctic Myrmica angulinodis, M. eidmanni, M. koreana, M. transsibirica, Plagiolepis manczshurica, Proformica buddhaensis, P. jacoti, Temnothorax kaszabi, T. mongolicus Camponotus tashcumiri, C. turkestanus, Cardiocondyla koshewnikovi, 15 Central Asian and Formica clarissima, F. orangea, F. subpilosa, Lasius przewalskii, Iran-Turanian Messor excursionis, Proformica coriacea, P. kaszabi, Tapinoma orthocephalum, Temnothorax melleus, Tetramorium armatum, T. concaviceps, T. inerme Formica pressilabris 1 Europe-Siberian Myrmica arnoldii, M. commarginata, M. pisarskii, Polyergus 4 Siberia-Mongolian nigerrimus Camponotus sachalinensis, C. saxatilis, Dolichoderus sibiricus, 10 East Asian Lasius fuji, Messor aciculatus, Myrmica forcipata, M. kamtschatica, M. kasczenkoi, Proformica mongolica, Tapinoma sinense Harpagoxenus zaisanicus 1 Mongolian endemic
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species from 18 genera in the Ural region of the surrounding regions or countries, regardless Russia; Reznikova (2003) revealed 70 species of land area. In general, ant species diversity (18 genera) along a transect from taiga-forest to declines with latitude and altitude elsewhere desert in western Siberia and eastern Kazakhstan. (Kusnezov 1957; Cushman et al. 1993; Dunn et Another Asian country, Iran, which has a similar al. 2009a, b; Guénard & Dunn 2012), and this land area to Mongolia, supports a much higher effect may, to some extent, explain the relatively diversity of ants, according to Paknia et al. (2008, low diversity of ants in Mongolia as the country 2010), Radchenko & Paknia (2010), Firouzi lies at northern latitudes and relatively high et al. (2011), Ghahari & Collingwood (2011), altitudes possessing a cold climate. Mohammadi et al. (2012), and Nezhad et al. Concerning the faunal composition (2012), who altogether listed 166 species (33 of ants in Mongolia, two major subfamilies, genera) from this country. Kupyanskaya (1995) Formicinae and Myrmicinae, comprise more than reported 79 species (24 genera) in the Russian 95% of the total recorded species, while another Far East. Concerning northern China, Guénard subfamily, Dolichoderinae, is represented by only & Dunn (2012) reported 128 species (19 genera) three species. The Formicinae and Myrmicinae from Xinjiang, 26 species (10 genera) from Inner are the largest ant subfamilies in the world and the Mongolia, 26 species (24 genera) from Gansu, 32 dominant groups in most terrestrial habitats. The species (10 genera) from Heilongjiang, and 32 prevalence of these subfamilies has been reported species (14 genera) from Jilin provinces. Three to increase with increasing aridity (Marsh 1986; other East Asian countries, more distantly located Lindsey & Skinner 2001). In contrast, Australian from Mongolia, also have a much higher diversity and North American semi-arid areas are of ants: Kim & Park (2003) reported 136 species characterised by a more even spread of species (39 genera) from South Korea; Radchenko (2005) across subfamilies such as the Dolichoderinae and found 99 species (35 genera) in North Korea, and Ponerinae (Briese & Macauley 1977; Whitford et Imai et al. (2003) listed 275 species (56 genera) al. 1978; Andersen 1986). This may reflect the from Japan. different evolutionary histories of the ant faunas This might reflect the intensity of the of the arid areas in these places. investigations carried out in different countries Almost half of the recorded species in or regions. It should be noted that, with the the Mongolian ant fauna belong to the genera exception of Japan, the ant faunas of most of the Formica and Myrmica. Five other genera contain nearby regions (especially in northern China) four or more species in Mongolia: Camponotus have not been fully explored, and the number of (5 spp.), Lasius (7 spp.), Proformica (5 spp.), species known from those regions may increase Temnothorax (4 spp.) and Tetramorium (4 significantly when remote or unexplored areas spp.). Together, the seven most diverse genera have been surveyed and all available material constitute nearly 81% of the ant species known has been examined (see Guénard & Dunn 2012). from Mongolia, while other genera, such as Nevertheless, current information is considered Cardiocondyla, Cataglyphis, Crematogaster, to be adequate to gain an understanding of the Dolichoderus, Harpagoxenus, Leptothorax, major faunal patterns in these regions (see Kim & Messor, Plagiolepis, Polyergus and Tapinoma, Park 2003; Radchenko 2005; Paknia et al. 2008; comprise a much smaller proportion of the fauna. Guénard & Dunn 2012). Similar faunistic patterns are found in other On the other hand, the extremely harsh arid Asian regions, e.g. eastern Kazakhstan and and fluctuating climates, as well as arid or very western Siberia (Reznikova 2003), the Ural region dry conditions in most areas of Mongolia, make of Russia (Gridina 2003), Kyrgyzstan (Tarbinsky the country unfavourable for the habitation of 1989; Schultz et al. 2006) and northern China species that require humid, sub-continental (Guénard & Dunn 2012). mild climates. This pattern of climatic and We suppose that the main faunal environmental conditions in Mongolia might be composition of ants of Mongolia is already considered the main basis for the relatively poor revealed, but some cryptic species likely remain composition of the ant fauna as compared with undiscovered because previous collecting has
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been mostly limited to nest collections, pitfall and found that the steppe was dominated by cold- traps and rarely baited traps. Extraction of ants resistant species; semi-desert supported mainly from sifted leaf litter using Winkler traps has opportunistic species, whereas desert involved proved to be very productive when conducting mostly hot climate specialists. However, these systematic inventories of ants (Fisher 1999; authors, and also Morton & Davidson (1988) Longino et al. 2002), but – to our knowledge – and Medel (1995), found no correlation between still has not been conducted in higher altitudes. precipitation and diversity of ants. Generally, the species richness of Distribution in phytogeographical regions local ant communities is primarily influenced by climatic conditions (e.g. temperature Based on the similarity analysis of the ant regimes, precipitation), landform, nest sites, faunas of various phytogeographical regions, microhabitat patterns (e.g. vegetation cover, three main groups can be distinguished. The soil type, moisture, texture), and food resource “forest-steppe” group encompasses the Khentii, availability (Koen 1988; Andersen 1993, 2000; Mongolian Dauria, Khangai, Middle Khalkh, Bestelmeyer & Wiens 2001; Fergnani et al. Eastern Mongolia and Great Khyangan regions. 2008). In Mongolia, however, due to the high The “steppe-semidesert” group includes the habitat heterogeneity, species diversity of ants Khovd, Depression of Great Lakes, Mongolian in various phytogeographical regions may differ Altai, Valley of Lakes, Eastern Gobi, Gobi Altai as a consequence of small microclimatic factors and Trans-Altai Gobi regions. The “desert” group that can determine whether a species is present or consists of the Alashan Gobi and Dzungarian absent within a location (Pfeiffer et al. 2003); thus Gobi regions. Surprisingly, the Khuvsgul taiga small-scale patterns of habitat distribution are forest region fell within “steppe-semidesert” important for ant life in Mongolia, especially in group, but we consider this is because of the very arid regions. Li et al. (2011) revealed that in arid low ant diversity discovered in Khuvsgul region, ecosystems of Central Asia, the species richness as only seven species were recorded there. As and abundance (nest density) of ants are closely we noted above, due to lower sampling intensity associated with silt content, soil organic matter, and completeness in various phytogeographical nitrogen and soil moisture, as well as topsoil regions of Mongolia, the actual diversity of ants temperature. Similarly, in humid subtropical and may increase,when remote or unexplored areas dry tropical areas, habitat strongly influences the have been surveyed. composition and community structure of ants It is notable that the most common species (Lindsey & Skinner 2001; Calcaterra et al. 2010). (e.g. Cataglyphis aenescens, Formica aquilonia, F. candida, F. kozlovi, F. lemani, F. pisarskii, Relation between plant and ant richness Proformica mongolica) tended to be widely distributed across various phytogeographical Positive correlation between plant diversity and regions, but in contrast, the rare species diversity of ants is frequently reported (Andersen (Dolichoderus sibiricus, F. pressilabris, Lasius 1997; Paknia & Pfeiffer 2012), and this was in reginae, L. obscuratus, Messor excursionis, accordance with our results on the relationship Myrmica commarginata, M. jessensis, M. between the diversity of ants and that of vascular kamtschatica, M. ruginodis, M. transsibirica, plants in the various phytogeographical regions Tapinoma orthocephalum, T. sinense, T. melleus, of Mongolia. Other investigations showed that Tetramorium concaviceps) were highly restricted habitats with more complex vegetation structure in their distribution, generally occurring only and higher primary productivity support more in one region. However, this might also be a diverse and dense ant assemblages (Andersen sampling artefact as rare species are hard to find 1997; Kaspari et al. 2000; Sarty et al. 2006; and detect. Paknia & Pfeiffer 2012). Moreover, in arid Pfeiffer et al. (2003) noted the habitats of central and southern Mongolia, considerable change of ant communities along Pfeiffer et al. (2003) revealed that structures of the environmental gradient from steppe to desert, ant and plant communities were highly correlated.
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These authors also indicated that the ant diversity It is worth noting that this is the first pattern could be a result of the productivity of report on the elevational distribution of ants resources that is a consequence of soil texture, in Mongolia, and the indistinct pattern of ant small-scale topography and other mosaic-like distribution in various elevation steps might also local environmental variations. be caused by varying investigation intensities in different regions of the country. This emphasises Elevational distribution the importance of intensive biogeographic research to get more realistic elevational We suggest that the unclear differentiation of distribution maps. elevational distribution for the majority of ant species is due to local mosaic-like topographic Biogeography relief, as the vast majority of the country contains mountainous landscapes (about 85% of its area is The biogeographic composition of ants in over 1,000 m a.s.l.), the exception being the plain Mongolia confirms it as one of the representative grasslands in eastern Mongolia. Nevertheless, the parts of the Palaearctic ant fauna, as there are observed patterns conform to the mid-elevational neither Oriental nor Afrotropical species. Such rise in species richness recorded in temperate a pattern was predicted by Fellowes (2006) at (Sanders et al. 2003; Glaser 2006), subtropical the generic level. The majority of ant species in (Bharti et al. 2013) and tropical regions Mongolia seem to be widespread in the whole (Fisher 2004). At similar altitudes, the northern Palaearctic region, with the addition of a few Palaearctic, Europe-Siberian and Boreo-montane Holarctic elements. Thus, species which are species show a mid-elevational peak between 900 widely distributed in these areas comprise more and 1,800 m a.s.l. in the Eastern Alps (Glaser 2006). than half of the total species (56%). The mid-elevational peak in ant The other specific character of the diversity is sometimes attributed to the more Mongolian ant fauna is the presence of strong open and dry litter habitat conditions and prey western, northern and eastern components. resource availability at mid-elevations. Based on Nearly one fifth (21%) of Mongolian species their research in tropical montane forests, Brühl are distributed in various regions or countries in et al. (1999) and Sabu et al. (2008) observed the Eastern Palaearctic region, such as Siberia, that ground ant abundance is higher in more Russian Far East, north-eastern China, Korean open and drier conditions, as moist and humid Peninsula and Japan. The remaining species habitat conditions limit ants’ foraging ability (22%) seem to be distributed in the arid regions and reduce the time availability for foraging on of Central Asia. Although the number of species the litter floor. shared with surrounding regions was relatively The absence of ant species at small, the whole ant fauna of Mongolia does not elevations above 2,500 m a.s.l. might be due differ significantly from that of the neighbouring to low temperature and strong winds, both of regions, and the majority of species occurring in which would preclude their distribution at high Mongolia probably originated in neighbouring altitudes in Mongolia. As ants are thermophiles areas to the west, north and east. (Hölldobler & Wilson 1990), which react Only a single species, Harpagoxenus negatively to low mean annual temperatures, the zaisanicus, is still known only from Mongolia cold climate seems to be a major factor working since its discovery, and thus is considered against the diversification of certain ant genera endemic. This species has a very cryptic life in Mongolia (Pfeiffer et al. 2003). Sanders et al. style as a nest parasite (see Pisarki 1963), and (2003) revealed that, generally, species of the is known from the dry steppe habitats in central, subfamilies Dolichoderinae and Myrmicinae are western and north-western Mongolia. There more common at lower elevations than at higher is a possibility of finding this species in areas elevations, while species of Formicinae are neighbouring Mongolia, especially in Siberia, more common at high elevations. However, we eastern Kazakhstan or northern China, and its did not observe such a pattern in altitudinal ant endemic status cannot be confirmed until more distributions in Mongolia. detailed studies are performed in these areas.
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There are several species occurring in the partial support of this research. The second the surrounding regions which might possibly author would like to thank the Japan Society be found in Mongolia in the future. These are for the Promotion of Science for the continuous Myrmica schencki Viereck, 1903, currently support of his research. We are grateful to Graham known from Kyrgyzstan, Xinjiang of China and T. Reels for professional language editing, to the Russian Far East; Lasius umbratus (Nylander, editor John R. Fellowes and to three anonymous 1846), currently recorded from Heilongjiang and reviewers for their helpful comments. Xinjiang provinces of China, southern Siberia, the European part and Far East of Russia, North Korea, South Korea, Japan and Taiwan; REFERENCES Proformica epinotalis Kuznetsov-Ugamsky, 1927, currently found in Heilongjiang and Aibek U, 2011.To the conservation of ants (Formicidae) Xinjiang provinces of China, Kyrgyzstan, Chita in Mongolia. Proceedings of Scientific region, Tuva and Buryat republics, Far East and Conference on the Issues and Current European part of Russia; and Camponotus itoi Status of Insect Conservation of Mongolia. Ulaanbaatar, 25 – 26 (in Mongolian). Forel, 1912, distributed in Xinjiang province of Aibek U & Yamane Sk, 2009. Taxonomic review of China, Taiwan, North Korea, South Korea and the genus Camponotus (Hymenoptera, Japan. Formicidae, Formicinae) from Mongolia. It should be mentioned here that we Biogeography 11: 97 – 108. excluded the exotic species, Monomorium Aibek U & Yamane Sk, 2010. Discovery of the pharaonis (Linnaeus, 1758) from the species list of subgenera Austrolasius and Dendrolasius Mongolian ants. This species is distributed nearly of the ant genus Lasius (Hymenoptera, all over the world, without any biogeographical Formicidae) from Mongolia. Japanese significance. In Mongolia it has been found only Journal of Systematic Entomology 16: 197 – 202. in a human settlement, and not in natural habitats Aibek U, Sonomdagva C & Yamane Sk, 2006. A (Aibek 2011). preliminary survey on the species composition Finally, it should be noted that seven and nesting habitats of ants in the Bogdkhan ant species found in Mongolia are listed as Mountain region, North Central Mongolia. threatened species in the Red List of the ANeT newsletter 8: 11 – 15. International Union for Conservation of Nature. Andersen AN, 1986. Diversity, seasonality and Thus, Harpagoxenus zaisanicus, Lasius reginae community organization of ants at adjacent and Polyergus nigerrimus are categorised as heath and woodland sites in south-eastern Vulnerable species, whereas Formica aquilonia, Australia. Australian Journal of Zoology 34: 53 – 64. F. lugubris, F. pratensis and F. uralensis are Andersen AN, 1993. Ant communities in the Gulf treated as Near-threatened species (IUCN 2012). Region of Australia’s semi-arid tropics: In order to conserve Mongolia’s biodiversity, species composition, patterns of organisation, the development of conservation strategies and biogeography. Australian Journal of for maintaining threatened ant species and the Zoology 41: 399 – 414. assessment of their endangerment are required Andersen AN, 1997. Functional groups and patterns for the future. of organization in North American ant communities: a comparison with Australia. ACKNOWLEDGEMENTS Journal of Biogeography 24: 433 – 460. Andersen AN, 2000. A global ecology of rainforest ants: Functional groups in relation to We would like to thank two students of the environmental stress and disturbance. In: National University of Mongolia, Ms Batsaikhan Ants: Standard Methods for Measuring Enkhdelger and Ms Ganbaatar Khulan, for their and Monitoring Biodiversity (Agosti, D, technical assistance in data-collecting. The first Majer JD, Alonso LE & Schultz TR, eds), author would like to thank the Korea Foundation Smithonian Institution Press, Washington, of Advanced Studies and the Asia Research London, 25 – 34. Centre at the National University of Mongolia for
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Seifert B, 1992. A taxonomic revision of the Palaearctic Tarbinsky YS, 1989. Myrmecological Complexes of members of the ant subgenus Lasius s. str. Tien-Shan and Alai. Abstract of doctoral (Hymenoptera: Formicidae). Abhandlungen thesis. Institute of Evolutionary Morphology und Berichte des Naturkundemuseums and Ecology of Animals, Russian Academy Goerlitz 66(5): 1 – 66. of Sciences. Moscow, 35 pp. (in Russian). Seifert B, 2000. A taxonomic revision of the ant Ulziikhutag N, 1989. Survey of the Flora of Mongolia. subgenus Coptoformica Mueller, 1923 State Press, Ulaanbaatar, 208 pp. (in (Hymenoptera, Formicidae). Zoosystema 22: Mongolian). 517 – 568. Wetterer JK & Radchenko A, 2011. Worldwide spread of Seifert B, 2002. The ant genus Cardiocondyla (Insecta: the ruby ant, Myrmica rubra (Hymenoptera: Hymenoptera: Formicidae): A taxonomic Formicidae). Myrmecological News 14: 87 – revision of the C. elegans, C. bulgarica, 96. C. batesii, C. nuda, C. shuckardi, C. Whitford WG, Johnson P & Ramirez J, 1978, Structure stambuloffii, C. wroughtonii, C. emeryi, and and seasonal activity of Chihuahuan Desert C. minutior species groups. SO - Annalen des and communities. Insectes Sociaux 25: 79 – Naturhistorischen Museums in Wien Serie B 88. Botanik und Zoologie 104B: 203 – 338. Yamane Sk & Aibek U, 2007. Ants of Mongolia. Nature Seifert B, 2004. The “Black Bog Ant” Formica picea and Insects 42: 20 – 25 (in Japanese) Nylander 1846 - a species different from Yamane Sk & Aibek U, 2012. Distribution of Myrmica Formica candida Smith 1878 (Hymenoptera: species (Hymenoptera, Formicidae) in Formicidae). Myrmecological News 6: 29 – Mongolia. Japanese Journal of Systematic 38. Entomology 18: 165 – 179. Seifert B & Schultz R, 2009. A taxonomic revision Yunatov AA, 1950. Main Patterns of the Vegetation of the Formica rufibarbis Fabricius, Cover of the Mongolian People’s Republic. 1793 group (Hymenoptera: Formicidae). Proceedings of the Mongolian Commission, Myrmecological News 12: 255 – 272. 39, 233 pp. (in Russian). Stitz H, 1934. Schwedisch-chinesische wissen- schaftliche Expedition nach den nordwestlichen Provinzen Chinas, unter Leitung von Dr. Sven Hedin und Prof. Sü Ping-chang. Insekten gesammelt vom schwedischen Arzt der Expedition Dr. David Hummel 1927 – 1930. 25. Hymenoptera. 3. Formicidae. Arkiv för Zoologi 27A: 1 – 9.
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Appendix
Ant species and their distributions in different phytogeographical regions of Mongolia
No Species name KHU KHE KHA MD GKH KHO MA MKH EM DGL VL EG GA DZG TAG AG 1 Camponotus aterrimus Emery, 1895 1 1 1 1 1 0 0 0 1 1 0 0 0 0 0 0 2 C. sachalinensis Forel, 1904 1 1 1 1 0 1 1 0 0 1 0 0 0 0 0 0 3 C. saxatilis Ruzsky, 1895 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 4 C. tashcumiri Tarbinsky, 1976 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 5 C. turkestanus André, 1982 0 0 0 1 0 0 1 0 0 1 0 1 1 1 1 0 6 Cardiocondyla koshewnikovi Ruzsky, 1902 0 0 0 0 0 0 1 0 0 1 0 0 1 0 1 0 7 Cataglyphis aenescens (Nylander, 1849) 1 0 1 0 0 1 1 1 0 1 1 1 1 0 1 0 8 Crematogaster subdentata Mayr, 1877 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 1 9 Dolichoderus sibiricus Emery, 1889 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 10 Formica aquilonia Yarrow, 1955 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 11 F. candida Smith, 1878 1 1 1 1 0 1 1 1 1 1 1 1 1 0 0 0 12 F. clara Forel, 1886 0 0 0 0 0 0 1 1 0 0 0 0 1 0 1 0 13 F. clarissima Emery, 1925 0 1 1 1 1 0 0 1 0 1 0 1 1 0 0 0 14 F. cunicularia Latreille, 1798 0 0 0 1 0 0 1 0 0 0 0 0 1 1 1 0 15 F. exsecta Nylander, 1846 0 1 0 1 0 0 0 0 1 0 0 0 0 0 0 0 16 F. forsslundi Lohmander, 1949 0 1 1 1 0 0 0 1 1 1 0 0 0 0 0 0 17 F. kozlovi Dlussky, 1965 1 1 1 1 0 1 0 1 0 1 0 1 1 0 1 0 18 F. lemani Bondroit, 1917 0 1 1 1 0 1 1 0 0 1 1 0 1 1 0 0 19 F. lugubris Zetterstedt, 1838 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 20 F. manchu Wheeler, 1929 0 1 1 1 0 0 0 1 0 1 0 0 0 0 0 0 21 F. orangea Seifert & Schultz, 2009 0 0 0 0 0 0 1 1 0 0 0 0 0 0 1 0 22 F. pisarskii Dlussky, 1964 1 1 1 1 0 1 1 1 1 0 0 0 0 0 0 0 23 F. pratensis Retzius, 1783 0 0 0 0 0 0 1 0 0 0 0 0 0 1 0 0 24 F. pressilabris Nylander, 1846 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 25 F. sanguine Latreille, 1798 0 1 0 1 0 1 1 1 1 0 0 0 0 0 0 0 26 F. subpilosa Ruzsky, 1902 0 0 1 0 0 1 1 1 0 1 0 0 0 0 0 0 27 F. truncorum Fabricius, 1804 0 1 1 0 0 0 1 0 0 0 1 0 0 0 0 0 28 F. uralensis Ruzsky, 1895 0 1 1 1 0 0 1 1 1 0 0 0 0 0 0 0 29 Harpagoxenus zaisanicus Pisarski, 1963 0 1 1 0 0 0 0 0 0 0 0 0 0 1 0 0 30 Lasius distinguendus (Emery, 1916) 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 31 L. fuji Radchenko, 2005 0 1 0 0 1 0 0 0 0 0 0 0 0 0 0 0 32 L. gebaueri Seifert, 1992 0 1 1 1 0 0 0 1 0 0 0 1 1 0 0 0 33 L. niger (Linnaeus, 1758) 0 0 0 0 0 1 1 0 0 1 0 0 0 0 0 0 34 L. obscuratus Stitz, 1930 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 35 L. przewalskii Ruzsky, 1915 0 1 0 0 0 0 0 0 0 0 0 1 0 0 0 0 36 L. reginae Faber, 1967 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 37 Lepthothorax acervorum (Fabricius, 1793) 1 1 1 1 0 1 1 0 0 0 0 0 0 0 0 0
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38 L. muscorum (Nylander, 1846) 0 1 1 1 0 0 1 1 0 1 0 0 0 0 0 0 39 Messor aciculatus (Smith, 1874) 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 40 M. excursionis Ruzsky, 1905 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 41 Myrmica angulinodis Ruzsky, 1905 0 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 42 M. arnoldii Dlussky, 1963 0 1 1 1 0 0 0 0 0 0 0 0 0 0 0 0 43 M. commarginata Ruzsky, 1905 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 44 M. divergens Karavaiev, 1931 0 1 1 0 0 0 1 1 0 1 0 0 0 0 0 0 45 M. eidmanni Menozzi, 1930 0 1 0 1 0 0 1 0 0 0 0 0 0 0 0 0 46 M. forcipata Karavaiev, 1931 0 1 1 1 0 0 0 1 1 0 0 0 0 0 0 0 47 M. kamtschatica Kupyanskaya, 1986 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 48 M. kasczenkoi Ruzsky, 1905 0 1 1 1 0 1 0 1 0 1 0 1 0 0 0 0 49 M. koreana Elmes, Radchenko et Kim, 2001 0 0 0 1 0 0 0 1 1 0 0 1 0 0 0 0 50 M. lobicornis Nylander, 1846 0 0 0 0 0 0 0 1 1 0 0 1 0 0 0 0 51 M. pisarskii Radchenko, 1994 0 1 1 1 0 0 0 1 1 0 0 0 0 0 0 0 52 M. ruginodis Nylander, 1846 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 53 M. sulcinodis Nylander, 1846 0 1 0 1 0 0 0 0 0 0 0 0 0 0 0 0 54 M. transsibirica Radchenko, 1994 0 1 0 0 0 0 0 0 0 0 0 0 0 0 0 0 55 Plagiolepis manczshurica Ruzsky, 1905 0 0 0 1 0 0 0 1 0 0 0 1 1 0 1 0 56 Polyergus nigerrimus Marikovsky, 1963 0 0 0 1 0 0 0 0 1 0 0 0 0 0 0 0 57 Proformica buddhaensis Ruzsky, 1915 0 1 1 1 0 1 0 1 0 1 0 1 0 0 0 0 58 P. coriacea Kuznetsov-Ugamsky, 1927 0 0 0 0 0 0 0 0 0 1 0 0 0 0 1 0 59 P. jacoti (Wheeler, 1923) 0 0 0 1 0 0 0 1 0 0 0 1 1 0 0 0 60 P. kaszabi Dlussky, 1969 0 0 0 0 0 0 0 1 0 1 0 1 1 0 0 0 61 P. mongolica Emery, 1901 0 1 1 1 0 1 1 1 1 1 1 1 1 0 0 0 62 Tapinoma orthocephalum Stitz, 1934 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 63 T. sinense Emery, 1925 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 64 Temnothorax kaszabi (Pisarski, 1969) 0 1 1 1 0 1 0 0 0 1 0 0 0 0 0 0 65 T. melleus (Forel, 1904) 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 66 T. mongolicus (Pisarski, 1969) 0 1 1 1 0 0 0 1 1 1 0 1 0 0 0 0 67 T. nassonovi (Ruzsky, 1895) 0 1 1 1 0 0 0 1 1 0 0 0 0 0 0 0 68 Tetramorium armatum Santschi, 1927 0 0 0 0 0 0 0 0 0 0 0 1 1 0 1 0 69 T. concaviceps Bursakov, 1984 0 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 70 T. inerme Mayr, 1877 0 0 0 0 0 0 1 0 0 0 0 0 0 0 1 0 71 T. tsushimae Emery, 1925 0 0 0 0 0 1 1 1 0 1 1 1 1 1 1 1
Refer to “Study area” subsection for abbreviations of the names of phytogeographical regions.
ASIAN MYRMECOLOGY A Journal of the International Network for the Study of Asian Ants Communicating Editor: John R. Fellowes
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